A new species of Bembidion (Ecuadion) from Ecuador (Coleoptera, Carabidae, Bembidiini), with a key to members of the georgeballi species group

A peer-reviewed open-access journa I

ZooKeys 249: 51-60 (2012)

doi: 10.3897/zookeys.249.4 149 RESEARCH ARTICLE #7,00Ke y

www.zookeys.o rg Launched to accelerate biodiversity research

A new species of Bembidion (Ecuadion) from Ecuador
(Coleoptera, Carabidae, Bembidiini), with a key to
members of the georgeballi species group

David R. Maddison'*, Luca Toledano**

| Department of Zoology, Oregon State University, Corvallis, OR 97331, USA 2 Museo Civico di Storia Na-
turale di Verona, Lungadige Porta Vittoria 9, 37129 Verona, Italy

T urn:lsid:zoobank.org:author:075A5E9B-5581-457D-8D2F-0B5834CDE04D
* urn:lsid:zoobank. org:author: DB6B 10C0-5035-468B-9632-9C337D50668B

Corresponding author: David Maddison (david.maddison@science.oregonstate.edu)

Academic editor: 7’ Erwin | Received 21 October 2012 | Accepted 27 November 2012 | Published 10 December 2012
urn:lsid:zoobank.org:pub: DC4D8668-A94C-44BC-BOF5-BALAD 91 1D 143

Citation: Maddison DR, Toledano L (2012) A new species of Bembidion (Ecuadion) from Ecuador (Coleoptera,
Carabidae, Bembidiini), with a key to members of the georgeballi species group. ZooKeys 249: 51-60. doi: 10.3897/
zookeys.249.4149

Abstract

A new species of ground beetle, Bembidion ricei, is described from the Andes mountains of Ecuador east of
Quito. It belongs to the georgeballi species group of subgenus Ecuadion, and is most similar to B. georgeballi.
A key to the species of the group is provided.

Keywords
Bembidion, Trechinae, Bembidiini, DNA, morphology, taxonomy, systematics

Introduction

Ecuadion is a diverse subgenus of Bembidion restricted to higher elevations in South
and Central America (Erwin 1982; Moret and Toledano 2002; Toledano 2008; Vigna
Taglianti and Toledano 2008). Adult beetles range in length from 2.2 to 6.1 mm;
they are generally shades of brown, either uniform or with various patterns, a few spe-

Copyright David R. Maddison, Luca Toledano. This is an open access article distributed under the terms of the Creative Commons Attribution License
3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

52 David R. Maddison & Luca Toledano / ZooKeys 249: 51-60 (2012)

cies having metallic reflections. Unlike many other Bembidion, most Ecuadion are not
closely associated with shores of bodies of water; they inhabit the leaf litter of cloud
forests (e.g., B. georgeballi Yoledano, B. onorei Moret and Toledano, B. andersoni To-
ledano), or run on clay cliffs along roadsides (e.g., B. agonoides Vigna Taglianti and
Toledano, B. chimborazonum Bates, B. walterrossii Toledano), or inhabit open high-
elevation grasslands (e.g., B. humboldti Moret and Toledano, B. guamani Moret and
Toledano, B. chimborazonum Bates, B. cotopaxi Moret and Toledano).

In Toledano’s (2008) study on the Northern Andean fauna of Bembidion, he il-
lustrated (his Figure 21) a single female belonging to the georgeballi species group
from Rio Chalpi, Ecuador, noting that it was similar to B. georgeballi but that it may
represent a separate species; the decision about its status was postponed until males
could be discovered. In 2010, the senior author and colleagues collected specimens
of this form from the same region from which Toledano’s specimen originated. The
larger series and characteristics of the male genitalia indicate that this form is a species
distinct from B. georgeballi, and it is here described, and compared to other members

of the georgeballi group.

Methods

Several hundred specimens of Ecuadion were examined as part of this study, including
50 specimens of B. georgeballi and 16 specimens of B. ricei, n. sp. Specimens came from
or have been deposited in the collections listed below. Each collection’s listing begins
with the coden used in the text.

BMNH._ The Natural History Museum, London

CTVR Luca Toledano Collection, Verona, Italy

MNHN Muséum National d’ Histoire Naturelle, Paris

OSAC Oregon State Arthropod Collection, Oregon State University, Corvallis
QCAZ = Catholic University of Ecuador, Quito

USNM_ National Museum of Natural History, Smithsonian Institution, Washington

Methods of specimen preparation for morphological work, and terms used, are
given in Maddison (1993; 2008). Measurements for Apparent Body Length (ABL) are
from apex of the labrum to apex of the longer elytron.

Photographs of body parts were taken with a Leica Z6 and JVC KY-F75U camera.
For pronotal, elytral, and genitalic images, a stack of photographs at different focal
planes was taken using Microvision’s Cartograph software; these photographs were
then merged using the PMax procedure in Zerene Systems’s Zerene Stacker; the im-
ages thus potentially have some artifacts caused by the merging algorithm.

Sequences of 28S ribosomal DNA and cytochrome oxidase I genes were obtained
using the protocols given in Maddison (2012), and deposited in GenBank with acces-
sion numbers JX971116 and JX971117.

A new species of Bembidion (Ecuadion) from Ecuador (Coleoptera, Carabidae, Bembidiini)... 53

Common morphological features and composition of the georgeballi species group

Among Ecuadion, adults of the georgeballi group are characterized by convex elytral in-
tervals, with deep and complete elytral striae, and with elytra reddish or with yellowish
markings (Toledano 2008). There are five known species in the group, four of which
are restricted to Ecuador (Fig. 1), the fifth in Venezuela:

B. georgeballi Toledano.........sseseeees [Ecuador]
B. ricei Maddison and Toledano....... [Ecuador]
B. pierre Voledanonissn dead Aiecats, [Ecuador]
B. cosangaense Voledano .........:.10006+ [Ecuador]
B. guaramacal Toledano ...........04. [Venezuela]

Species identification and description

1 Elytra reddish, unicolorous; if with a faint, slightly darker spot, then with a
pale rufous pronotum (see Toledano 2008: Fig. 24) oo... eee eeeseeseeeeeeeeees 2
— Elytra with a mottled testaceous and brown pattern, pronotum darker, at least
CEDAR VStLIO. ents. retedee tutta settee ett ant te tM Lath pontbybicetgu: 3
2 Pronotum piceous-black, narrower (pronotal width/length = 1.17 to 1.20),
elytra reddish, unicolorous; EcuadOt. c:.ct tie cescevetsctevsece sevenaet B. cosangaense

= Pronotum rufous, wider (pronotal width/length = 1.32 to 1.33), elytra red-
dish, sometimes with a faint, slightly darker spot; Venezuela... B. guaramacal

3 Posterior lateral seta and carina of pronotum both absent, pronotum more
constricted-atdhind tarigles (Pig OB )e csve.ckedhdande. guvSebeges asics B. georgeballi
- Posterior lateral seta and carina of pronotum both present, pronotum less
constricted athindsang less (eles, Fite P WN) exec necnensvhovader tomes sh savai smelt nasty ttehaie 4
= Microsculpture absent from elytra in males and females; elytral striae 3 and
4 connected in front of the anterior discal seta (see Fig. 2A and 4A); elytral
PLES Val SAIN Ota By MOOT VERE tats cect ore oh sen tie a leeneatet eee B. ricei sp. n.

a Isodiametric microsculpture on the whole dorsal surface in males and fe-
males; elytral striae normal; elytral intervals only slightly convex....B. pierrei

Bembidion ricei Maddison & Toledano, sp. n.
urn:lsid:zoobank.org:act:A7 2AEO D9-3325-427C-B59B-E0B02507CA9B
http://species-id.net/wiki/Bembidion_ricei

Holotype. Adult male, with three labels: “ECUADOR: Napo: Rio Chalpi Grande,
2800m, 0.3645°S 78.0852°W, 26.x.2010. DRM 10.159. W.P. & D.R. Maddison, M.
Reyes”, “David R. Maddison DNA2653 DNA Voucher [printed on pale green paper]”,
and “HOLOTYPE Bembidion ricei Maddison & Toledano [printed on red paper]”.

54 David R. Maddison & Luca Toledano / ZooKeys 249: 51-60 (2012)

® B. ricei
@ B. georgeballi
m@ B. cosangense
A B. pierrei

Figure |. Geographic distribution of known members of the georgeballi species group in Ecuador. Base
map modified from Chirico and Warner (2005).

Specimen to be deposited at QCAZ; temporarily in OSAC. Genitalia in glycerine in
vial pinned beneath specimen. GenBank accession numbers of DNA sequences from
the holotype: JX971116 (28S ribosomal DNA) and JX971117 (cytochrome oxidase I).

Paratypes. Six males and nine females from: ECUADOR: Napo: Rio Chalpi Grande,
2800m, 0.3645°S, 78.0852°W, 26.x.2010 & 8.xi.2010 (11 exx., OSAC, BMNH,
MNHN); ECUADOR: Napo: Papallacta, 2750 m, Rio Chalpi, 8.xi.1985 (1 ex., CTVR);
ECUADOR: Napo: Rio Guango (=Rio Huango), 2730m, 0.3758°S, 78.0748°W,
26.x.2010 (2 exx., OSAC); ECUADOR: Napo: Sierrazul (Hacienda Aragon) 10 km W
of Cosanga, 2250m, 3—4.vi.1993, 00°44’08"S, 077°53’50"W (1 ex., USNM).

Type locality. ECUADOR: Napo: Rio Chalpi Grande, 2800m, 0.3645°S,
78.0852°W.

A new species of Bembidion (Ecuadion) from Ecuador (Coleoptera, Carabidae, Bembidiini).... 55

Figure 2. Habitus of male B. ricei and B. georgeballi. Scale bar is 1 mm. A B. ricei (ECUADOR: Napo:
Rio Chalpi Grande, 2800m, 0.3645°S, 78.0852°W, D.R. Maddison voucher V100622) B B. georgeballi
(ECUADOR: Pichincha: Quebrada Lozada, on road to Res. Yanacocha, 3460m, 0.1105°S, 78.5642°W,
voucher V100658).

Derivation of specific epithet. It gives us great pleasure to name this species after
the late Harold Edward Rice, a passionate butterfly collector and active member of the
Pacific Northwest lepidopterist community, and friend to the senior author. Through
Harold’s generosity, systematic entomology is well supported at Oregon State Univer-
sity. The fund he established paid for the expedition that yielded most of the known
specimens of B. ricei, including the holotype.

Diagnosis. A shiny, medium-sized Bembidion (Ecuadion) with convex elytral in-
tervals, and with a mottled pattern of light and dark on the elytra (as in Fig. 2A); adults
have a seta at the hind corner of the pronotum, and lack elytral microsculpture in both
males and females. Stria 3 and stria 4 are joined together and interrupted in front of
the anterior discal seta (ed3; Fig. 2A, 4A); the striae are otherwise complete, and deeply
engraved. This combination of characteristics is distinct within the genus.

Brown, with lateral margins of pronotum paler brown in most specimens, and
with elytra having a pale apex, and a pale transverse preapical region surrounded by
darker brown (Fig. 2A); the region adjacent to ed3 is also slightly darker. Prothorax
with sinuate lateral margin, hind angles about 90°, and with a posterior lateral carina
(Fig. 3A). Elytral striae deep, complete, although with striae 3 and 4 joined together
and interrupted in front of ed3 (Fig. 4A) in all 16 specimens examined; elytral intervals

56 David R. Maddison & Luca Toledano / ZooKeys 249: 51-60 (2012)

. -. a io es ea .
Figure 3. Pronota of male B. ricei and B. georgeballi. Scale bar is 0.1 mm. A B. ricei (ECUADOR: Napo:
Rio Chalpi Grande, 2800m, 0.3645°S 78.0852°W, D.R. Maddison voucher V100677) B B. georgeballi
(ECUADOR: Pichincha: Quebrada Lozada, on road to Res. Yanacocha, 3460m, 0.1105°S, 78.5642°W,
voucher V100658).

Figure 4. Elytra of B. ricei (A & C, from ECUADOR: Napo: Rio Chalpi Grande, 2800m, 0.3645°S,
78.0852°W) and B. georgeballi (B & D, from ECUADOR: Pichincha: Quebrada Lozada, on road to Res.
Yanacocha, 3460m, 0.1105°S 78.5642°W). Scale bar is 0.1 mm. A B. ricei male (D.R. Maddison voucher
V100656), showing joining of striae 3 and 4 (indicated by arrows), with subsequent gap in each stria

(between arrows); illuminated by two diffuse lateral lights B B. georgeballi female (voucher V100676);
illuminated by two diffuse lateral lights C B. ricei female (voucher V100675), illuminated by a ring light
D B. georgeballi female (voucher V100676), illuminated by a ring light.

A new species of Bembidion (Ecuadion) from Ecuador (Coleoptera, Carabidae, Bembidiini).... 57

Figure 5. Male aedeagus. Scale bar is 0.1 mm. A B. ricei (ECUADOR: Napo: Rio Chalpi Grande,
2800m, 0.3645°S, 78.0852°W, D.R. Maddison voucher V100656) B B. georgeballi (ECUADOR:
Pichincha: Campamento Pichan, 3350m, 0.1093°S 78.5728°W, D.R. Maddison voucher V100657).

convex. Microsculpture absent from the pronotum and elytra in both sexes. Aedeagus
as in Fig. 5A. ABL 3.9-4.1mm, n=6.

One of the more unusual aspects of these Bembidion, shared with some other
Ecuadion, including males of B. georgeballi, is the amber-like clarity of the elytra. The
elytra are similar to clear lacquer in places, allowing bright microsculpture dots from
the undersurface of the elytra to be visible dorsally (see pale dots in Fig. 4C).

The only other Ecuadion with yellow and brown mottled elytra, and with complete
striae and convex elytral intervals, are B. georgeballi and B. pierrei (Toledano 2008).
B. ricei is most similar to B. georgeballi, sharing more convex elytral intervals, and pale
lateral regions of the pronotum in most specimens. Specimens differ in being larger
(ABL 3.9-4.1mm in B. ricei, 2.9-3.4mm in B. georgeballi), with a less constricted
posterior margin of the pronotum (compare Fig. 3A to 3B). The posterior lateral seta

58 David R. Maddison & Luca Toledano / ZooKeys 249: 51-60 (2012)

Figure 6. Type locality of Bembidion ricei, at Ecuador: Napo: Rio Chalpi Grande, 2800m, 0.3645°S,
78.0852°W. Specimens were found in damp leaf litter under rocks that had previously been at the site
marked by the arrow; this is a small tributary of the Rio Chalpi Grande, within 4m of that river. Several
Andinodontis muellermotzfeldi Toledano and Erwin were found within a meter of the B. ricei habitat; B.
(Ecuadion) sanctaemarthae Darlington was common about 2—4m away, along the upper banks of the main
river. One of the other known localities was along the same small creek, but upstream, in a more shaded

area, and further away from the shore (about 1—2m from the water), among damp leaf litter and rocks.

and carina of the pronotum are present (absent in B. georgeballi). B. ricei specimens
have striae 3 and 4 joined and interrupted in front of discal seta ed3 (Fig. 4A). Micro-
sculpture is lacking from the dorsal surface of the elytra in both males and females of
B. ricei, and thus they are very shiny (Fig. 4A, C); in B. georgeballi, males lack elytral
microsculpture, but females have evident isodiametric microsculpture throughout the
elytra (Fig. 4B, D). In addition, the male aedeagus of B. ricei has larger and darker
sclerotized regions on the internal sac (Fig. 5A). B. georgeballi is currently known only
from 3350—3550m on the slopes of Volcan Guagua Pichincha west of Quito (Fig. 1).

From B. pierrei, B. ricei can be distinguished by having striae 3 and 4 joined,
and lacking microsculpture on the elytra (B. pierrei has isodiametric microsculpture
throughout the elytra in both males and females). B. pierrei also lacks the transparent,
lacquer-like elytral regions of B. ricei. B. pierrei is known from the province of Chim-
borazo, far south of the localities of known localities of B. ricei (Fig. 1).

A new species of Bembidion (Ecuadion) from Ecuador (Coleoptera, Carabidae, Bembidiini)..._ 59

Geographic distribution. B. ricei occurs in the province of Napo between 2250m
and 2800m in the Andes of Ecuador, east of Quito (Fig. 1). Most specimens have been
found a few kilometers east of Papallacta along two tributaries of the Rio Papallacta; a
single female has been found along a tributary of the Rio Jondachi south of Cosanga
and west of La Merced de Jondachi.

Habitat. Found among leaf litter and under rocks in moist areas near small streams
in montane forest (Fig. 6). Specimens were found during daytime in leaf litter under
rocks or by scratching open leaf litter.

Acknowledgements

We thank Terry Erwin (USNM) for lending to us one of the paratypes. We are grate-
ful to Pierre Moret (Toulouse, France) who kindly donated to L.T. the paratype in
CTVR. We also thank David Kavanaugh and an anonymous reviewer for their valu-
able comments on the manuscript.

We are very thankful to all those who helped with a collecting expedition to Ec-
uador during which most of the type series of B. ricei and numerous other Ecuadion
were collected. Mauricio Vega arranged many details of the trip, including the rel-
evant collecting and export permits; Wayne Maddison, Marco Reyes, and Mauricio
Vega accompanied the senior author into the field and helped collect the specimens.

This project was funded by the Harold E. and Leona M. Rice Endowment at Or-
egon State University.

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